Seasonal changes in the response of oystercatchers Haematopus ostralegus to human disturbance

The response of foraging animals to human disturbance can be considered as a trade-off between the increased perceived predation risk of tolerating disturbance and the increased starvation risk of not feeding and avoiding disturbance. We show how the response of overwintering oystercatchers Haematopus ostralegus to disturbance is related to their starvation risk of avoiding disturbance. As winter progresses, oystercatcher energy requirements increase and their feeding conditions deteriorate. To survive they spend longer feeding and so have less spare time in which to compensate for disturbance. Later in winter, birds approach a disturbance source more closely and return more quickly after a disturbance. Their behavioural response to disturbance is less when they are having more difficulty surviving and hence their starvation risk of avoiding disturbance is greater. These results have implications for studies which assume that a larger behavioural response means that a species is more vulnerable to disturbance. The opposite may be true. To more fully understand the impact of disturbance, studies should measure both behavioural responses and the ease with which animals are meeting their requirements. Conservation effort should be directed towards species which need to spend a high proportion of their time feeding, but still have a large response to disturbance.     

Experimental evidence for a causal effect of pair-bond duration on reproductive performance in oystercatchers (Haematopus ostralegus)

Many studies have suggested that reproductive performance improvesduring the pair-bond, which might explain why individuals rematewith the same partner in many species. However, discussion existsabout whether the association between reproductive performanceand pair-bond duration that is reported in these studies reflectsa causal relationship. Usually it is unclear whether a positiveassociation is caused by pairs improving during their pair-bondor by high-quality pairs staying together for longer. Furthermore,reproductive performance often also depends on the age or breedingexperience of parents, which all covary with pair-bond duration.A much needed experimental approach is lacking so far. We investigatedthe effect of pair-bond duration on reproductive performancein a long-lived monogamous bird species based on natural aswell as experimental variation. The duration of oystercatcher(Haematopus ostralegus) pair-bonds, which were followed for21 years, strongly affected reproductive output, even aftercontrolling for effects of age and breeding experience. Pairsimproved during their pair-bond, and there were no indicationsof selective disappearance of low-quality pairs; however, pairsthat stayed together for very long performed badly. Experimentalremoval of one partner showed that the reproductive cost ofdivorce depended on the pair-bond duration with the old partner.In addition, after remating, the newly formed pairs stronglyimproved again, independent of the age and breeding experienceof the remated pair members. As such, this study provides thefirst experimental evidence of a causal effect of pair-bondduration on reproductive performance.     

Genetic analysis of the population structure of socially organized oystercatchers (Haematopus ostralegus) using microsatellites

On the island of Schiermonnikoog (The Netherlands), the breeding population of oystercatchers can be divided into two groups: 'residents' and 'leapfrogs', based on their distinct social characteristics and limited probabilities of status change between breeding seasons. In order to investigate whether this social organization has caused local genetic differentiation, leapfrogs and residents were compared at eight polymorphic microsatellite loci. No significant genetic subdivision between residents and leapfrogs was observed (theta = 0.0000; 95% confidence interval (CI), -0.0027-0.0033), indicating that the oystercatcher population on the island of Schiermonnikoog has to be considered as one panmictic unit. Investigation of three additional locations in the northern part of The Netherlands did not reveal significant genetic population subdivision either (theta = -0.0005; 95% CI, -0.0045-0.0037), despite the fact that adult osytercatchers show extreme fidelity to their breeding localities. These results indicate panmixis and considerable levels of gene flow within the northern part of The Netherlands. Thus, the results from genetical analyses do not seem to be in agreement with observational data on the dispersal behaviour of breeding individuals. It is argued that the lack of population structure, locally on Schiermonnikoog as well as across larger geographical distances, is to be attributed to high levels of gene flow through dispersal of juvenile birds.     

Field tests of the accuracy of estimating prey size from bill length in oystercatchers,Haematopus ostralegus,eating mussels,Mytilus edulis

Field tests were made of the accuracy with which different observers estimated the size of mussels, Mytilus edulis, being eaten by oystercatchers, Haematopus ostralegus. Mussel size was estimated by eye by comparing their lengths with that of the bird's bill. Tests of bias were made in two ways. A ‘natural’ test involved comparing the estimated sizes of prey taken by free-living birds with the lengths of the emptied shells they left on the mussel bed. A ‘model’ test involved estimating mussel size against the bill of a model oystercatcher. Though one observer showed no bias, the other three overestimated the lengths of large mussels and underestimated those of small ones. The magnitude of the bias differed significantly between observers and, to a lesser degree, between the natural and model tests. Because of the curvilinear relationship between prey length and energy content, a bias of the magnitude found in this study could, in extreme cases, lead to serious errors in estimating a prey's energy content. It is concluded that, in studies where prey size is measured against bill length, an observer's individual bias should always be measured, preferably by the natural method. If this is not possible, then tests of the robustness of the conclusions should be made by recalculating the data assuming worst case errors.     

Cerastoderma glaucum 5S ribosomal DNA: characterization of the repeat unit, divergence with respect to Cerastoderma edule, and PCR-RFLPs for the identification of both cockles.

The 5S rDNA repeat unit of the cockle Cerastoderma glaucum from the Mediterranean and Baltic coasts was PCR amplified and sequenced. The length of the units was 539-568 bp, of which 120 bp were assigned to the 5S rRNA gene and 419-448 bp to the spacer region, and the G/C content was 46%-49%, 54%, and 44%-47%, respectively. Two types of units (A and B), differing in the spacer, were distinguished based on the percentage of differences and clustering in phylogenetic trees. A PCR assay with specific primers for each unit type indicated that the occurrence of both units is not restricted to the sequenced individuals. The 5S rDNA units of C. glaucum were compared with new and previously reported sequences of Cerastoderma edule. The degree of variation observed in C. edule was lower than that in C. glaucum and evidence for the existence of units A and B in C. edule was not found. The two cockles have the same coding region but displayed numerous fixed differences in the spacer region and group separately in the phylogenetic trees. Digestion of the 5S rDNA PCR product with the restriction enzymes HaeIII and EcoRV revealed two RFLPs useful for cockle identification.     

Predation of cockles (Cerastoderma edule) by the whelk (Buccinum undatum) under laboratory conditions

The feeding rate and behaviour of whelks (Buccinum undatum)offered cockles (Cerastoderma edule) in laboratory experimentswere examined. When presented with cockles in a range of sizes(10–40 mm), 14 B. undatum (34.6–88.3 mm),held individually in aquaria, consumed a wide size range ofcockles. Small whelks (<40 mm) consumed cockles (<23 mm),whereas large whelks, (>60 mm) ate a greater numberof larger cockles (>30 mm) and a wider size range ofcockles (12–40 mm) than smaller whelks. The majority(90%) of the shells of the predated cockles were undamaged andthe few (<10%) that were damaged showed only slight abrasionsto the anterior and posterior shell margin. Filmed observationsof B. undatum feeding on C. edule showed a method of attackthat has not previously been reported and involved the use ofthe whelk's foot to asphyxiate the cockle or to pull the shellvalves apart. No filmed evidence was found for the previouslyreported shell ‘wedging’ technique for prising openthe closed shell valves of C. edule, although 10% of the shellsof consumed cockles in feeding experiments had damaged shellmargins. (Received 4 April 2007; accepted 30 June 2007)     

A silver spoon for a golden future: long-term effects of natal origin on fitness prospects of oystercatchers (Haematopus ostralegus)

1. Long-term effects of conditions during early development on fitness are important for life history evolution and population ecology. Using multistrata mark-recapture models on 20 years of data, we quantified the relation between rearing conditions and lifetime fitness in a long-lived shorebird, the oystercatcher (Haematopus ostralegus). We addressed specifically the relative contribution of short- and long-term effects of rearing conditions to overall fitness consequences. 2. Rearing conditions were defined by differences in natal habitat quality, in which there is a clear dichotomy in our study population. In the first year of life, fledglings from high-quality natal origin had a 1.3 times higher juvenile survival. Later in life (age 3-11), individuals of high-quality natal origin had a 1.6 times higher adult prebreeder survival. The most striking effect of natal habitat quality was that birds that were reared on high-quality territories had a higher probability of settling in high-quality habitat (44% vs. 6%). Lifetime reproductive success of individuals born in high-quality habitat was 2.2 times higher than that of individuals born in low-quality habitat. This difference increased further when fitness was calculated over several generations, due to a correlation between the quality of rearing conditions of parents and their offspring. 3. Long-term effects of early conditions contributed more to overall fitness differences as short-term consequences, contrary to common conceptions on this issue. 4. This study illustrates that investigating only short-term effects of early conditions can lead to the large underestimation of fitness consequences. We discuss how long-term consequences of early conditions may affect settlement decisions and source-sink population interactions.     

The palearctic oystercatcher Haematopus ostralegus, a natural definitive host for Gymnophalloides seoi

In order to verify the role of migrating birds as natural definitive hosts for Gymnophalloides seoi (Gymnophallidae), Palearctic oystercatchers, Haematopus ostralegus, were caught from several western coastal areas in the Republic of Korea and examined for intestinal flukes. Five (71.4%) of 7 oystercatchers were infected with 302-1,660 (mean 892) adult G. seoi. In intestinal sections of the host, worms were found in the intervillous space of the mucosal layer. We conclude that the oystercatcher is a natural definitive host for G. seoi.     

The effect of size and temperature on the predation of cockles Cerastoderma edule (L.) by the shore crab Carcinus maenas (L.)

Laboratory studies demonstrated that shore crabs Carcinus maenas (L.) can consume <40 cockles Cerastoderma (= Cardium) edule (L.) per individual · day⁻¹. Various predation techniques used by the crabs are reported. The time required to open and consume individual cockles increased exponentially with prey size. Small (<l5-mm shell height), easily broken cockles appeared to be the most profitable in terms of energy acquisition per unit of handling time, the optimal size of prey increasing with predator size. With unlimited prey available, however, crabs selected prey of mean size smaller than these predicted optima, and much below the maximum size they were capable of opening. Feeding rates, both in terms of cockles ingested or energy intake per day, rose steeply with increasing temperature, but the size range of prey consumed remained unchanged. These data strongly suggest that Carcinus maenas is a potentially important predator of small cockles, particularly during the wanner summer months.     

The pair bond and divorce among Oystercatchers Haematopus ostralegus on Skokholm Island, Wales

Mate changes of Oystercatchers were followed over a 15-year period. In 8.0% of instances, the pair split up between breeding seasons even when both members bred the second season. The likelihood of divorce was highest for birds aged five years, the modal age of first breeding. After divorce birds did not select an older or younger mate when re-pairing. Low hatching success increased the chances of divorce. Divorce was not followed by significant changes in clutch-size, laying date or nesting success (although females tended to be more successful).     

The cranial and cervical osteology of the european oystercatcher Haematopus ostralegus L.

The cranial and cervical osteology of the European oystercatcher, Haematopus ostralegus L. is completely described from both whole skeletons and spirit specimens. Contrary to previous reports, the postorbital ligament shows variability in bifurcation and attachment, with the posterior branch, where present, attaching either to the suprameatic process or the zygomatic process. In addition, the quadrate ligament, which had been described as originating from the suprameatic process, appears instead to attach to the zygomatic process in some specimens. The caudal mandibular fenestra, earlier considered absent in oystercatchers, is present in all specimens examined. The cranium is additionally distinguished by a number of features which may be unique to oystercatchers. A fourth, previously unrecorded, division of the maxillopalatine strut is present, while the palatine process of the premaxilla is reduced to a thickening along the edge of the premaxillary process of the palatine. The distinctiveness of the oystercatcher cranium is particularly evident in the area of the quadratomandibular articulation, which possesses features potentially important for stabilizing the joint. A deep retroarticular notch is present, guiding a large medial jugomandibular ligament along the posterior margin of the articulation. A lateral mandibular tuberosity, which is received dorsally by an emargination of the jugal arch and quadrate, may act as an osteological brace, preventing posterior shifting of the closed mandible. The 15 cervical vertebrae are divided on the basis of structural criteria into three sections.     

A proliferative disorder of possible hemic origin in the common cockle,Cerastoderma edule

A study is described concerning the infestation of a laboratory-reared Daphnia magna culture by the fungus Aphanomyces daphniae. The origin of this fungus is Lake Huron, Michigan, the water from which is used in the laboratory to culture daphnids and maintain fishes for toxicological testing. Infestations originate from the Lake during the autumnal and vernal homothermal periods. The fungus is apparently maintained in the laboratory in conjunction with the fishes. Onset of fungal infestations in zooplankton culture apparently is initiated by three types of stress: rising temperatures, declining dissolved oxygen values, and poor nutrition. These infections have been eliminated by separating the daphnid culturing facility from the fish facility, substituting an algal diet for a synthetic one, sterilizing all water, air, and glassware connected with zooplankton culture, and maintaining the daphnids in a constant temperature room with controlled dissolved oxygen levels in the culturing vessels.     

Individual and age differences in the feeding ecology of Oystercatchers Haematopus ostralegus wintering on the Exe Estuary, Devon

The habitats and foods used by a sample of marked individuals and by the whole population of Oystercatchers Haematopus ostralegus wintering on the Exe estuary, South Devon, are described. In the spring and early summer, only a few hundred immatures were present. Two thousand adults and several hundred juveniles arrived in late summer from the breeding grounds and remained until the following spring. The majority of the birds were then adults. At low water, most adults fed on the mussel Mytilus edulis beds and ate mussels. Most individuals specialised on this prey and ate little else. Some adults fed on mud-flats and sandflats within the estuary and along the coast, and specialised on a mixture of Nereis diversicolor and Scrobicularia plana or on Cerastoderma edule or Littorina spp. Though some juveniles ate mussels from the time they arrived, most did not. In winter, they took Scrobicularia on mudflats, earthworms Lumbricidae and leatherjackets Tipulidae larvae in fields and Spisula and mussels along the coast. At other times of year they mainly ate Nereis. However, as they grew older, more birds began to specialise on mussels, especially in their second, third and fourth summers when the adults were away. At high water, most birds roosted at the mouth of the estuary or along the coast. However, several hundred fed in the fields in winter with more doing so late in the winter on warm days on Neap tides when less time was available for feeding on the estuary in daylight. Birds feeding on Nereis and Scrobicularia at low water were most often seen in the fields, irrespective of age. Birds feeding on Mytilus and Littorina occurred there rarely, but no bird eating Cerastoderma was seen there. The findings are discussed in relation to age differences in feeding skills and ability to compete successfully with other birds.     

Modelling the population consequences of age- and sex-related differences in winter mortality in the oystercatcher, Haematopus ostralegus

A modelling approach is used to explore the effect of age and sex differences in oystercatcher ( Haematopus ostralegus ) winter mortality on population size, population structure and the population response to habitat loss or change. Increasing the mortality of first and second year birds reduced population size, but had very little effect on the proportion of the population that were adults. Increasing female mortality reduced population size and resulted in a male-biased population. A sex bias amongst birds of breeding age meant that there were fewer potential breeding pairs for a given population size, reducing the size of the breeding population and the breeding output. Increasing the mortality of one sex relative to the other reduced population size, even when mean adult mortality rates remained unchanged. Increasing the strength of density-dependent mortality in young birds caused a greater reduction in population size as habitat was lost. Increasing the strength of female density-dependent mortality had the same effect, even though male density-dependent mortality had been correspondingly reduced. Increasing density-independent or density-dependent winter mortality in one sex relative to another also exaggerated the disproportional effect of winter habitat loss on separate breeding subpopulations using the same overwintering area. These results suggest that any study of population dynamics should be aware of both age and sex differences in mortality. Conservationists should be particularly aware of any age or sex differences in diet or habitat use that may result in a differential response to environmental change.     

Influence of disseminated neoplasia, trematode infections and gametogenesis on surfacing and mortality in the cockle Cerastoderma edule

Cerastoderma edule is a widely distributed bivalve mollusc, commercially exploited throughout Europe and is also an important food source for birds and crustaceans. Recently, mass surfacing and mortalities of cockles have been observed and reported at sites in Ireland and elsewhere, particularly in the summer months. One such site is Flaxfort Strand, Courtmacsherry Bay, County Cork, Ireland, an important feeding area used by many seabirds during the summer months. For the past few years large numbers of surfaced cockles have been observed at the site in a moribund condition. Samples of cockles from this area were collected over the summer months and their health status assessed. Cockles that had surfaced (moribund) and those still buried in the sediment were quantified and screened: sex, gonadal maturity and size class of cockles were also determined. Disseminated neoplasia and trematodes were observed in screened cockles. The most significant finding during the study was that mortalities and surfacing of cockles was related to a greater incidence of disseminated neoplasia. No neoplasia was observed in the smallest and largest size classes. There was a significantly higher prevalence of neoplasia in moribund cockles than in buried cockles, whereas in both groups a similar concentration of trematode metacercariae was observed in the screened tissues. Also, most of the cockles that had surfaced were either in the process of spawning or were spent. Overall a much larger percentage of moribund cockles exhibited both trematode infections plus neoplasia compared with buried cockles. A combination of the presence of neoplasia and trematodes, along with stress related to spawning, may immunocompromise the cockless, causing the animals to surface and become moribund.